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We study optimal two-sector (vegetative and reproductive) allocation models of annual plants in temporally variable environments, that incorporate effects of density dependent lifetime variability and juvenile mortality in a fitness function whose expected value is maximized. Only special cases of arithmetic and geometric mean maximizers have previously been considered in the literature, and we also allow a wider range of production functions with diminishing returns. The model predicts that the time of maturity is pushed to an earlier date as the correlation between individual lifetimes increases, and while optimal schedules are bang-bang at the extremes, the transition is mediated by schedules where vegetative growth is mixed with reproduction for a wide intermediate range. The mixed growth lasts longer when the production function is less concave allowing for better leveraging of plant size when generating seeds. Analytic estimates are obtained for the power means that interpolate between arithmetic and geometric mean and correspond to partially correlated lifetime distributions., 29 pages, 12 figures

PREMISE The xylem tissue of plants performs three principal functions: transport of water, support of the plant body, and nutrient storage. Tradeoffs may arise because different structural requirements are associated with different functions or because suites of traits are under selection that relate to resource acquisition, use, and turnover. The structural and functional basis of xylem storage is not well established. We hypothesized that greater starch storage would be associated with greater sapwood parenchyma and reduced fibers, which would compromise resistance to xylem tensions during dehydration. METHODS We measured cavitation resistance, minimum water potential, starch content, and sapwood parenchyma and fiber area in 30 species of southern California chaparral shrubs (evergreen and deciduous). RESULTS We found that species storing greater starch within their xylem tended to avoid dehydration and were less cavitation resistant, and this was supported by phylogenetic independent contrasts. Greater sapwood starch was associated with greater parenchyma area and reduced fiber area. For species without living fibers, the associations with parenchyma were stronger, suggesting that living fibers may expand starch storage capacity while also contributing to the support function of the vascular tissue. Drought-deciduous species were associated with greater dehydration avoidance than evergreens. CONCLUSIONS Evolutionary forces have led to an association between starch storage and dehydration resistance as part of an adaptive suite of traits. We found evidence for a tradeoff between tissue mechanical traits and starch storage; moreover, the evolution of novel strategies, such as starch-storing living fibers, may mitigate the strength of this tradeoff.

Significance Plant vascular systems play a central role in global water and carbon cycles and drought resistance. These vascular systems perform multiple functions that affect the fitness of plants, and trade-offs are present among these functions. Some trade-offs are well established, but studies have not examined the full suite of functions of these complex systems. Here, we used a powerful multivariate method, structural equation modeling, to test hypotheses about the trade-offs that govern this vital and globally important tissue. We show that xylem traits are broadly governed by trade-offs related to transport, mechanical support, and storage, which are rooted in cellular structure, and that the level of dehydration experienced by plants in the field exerts a strong influence over these relationships., The xylem in plants is specialized to transport water, mechanically support the plant body, and store water and carbohydrates. Balancing these functions leads to trade-offs that are linked to xylem structure. We proposed a multivariate hypothesis regarding the main xylem functions and tested it using structural equation modeling. We sampled 29 native shrub species from field sites in semiarid Southern California. We quantified xylem water transport (embolism resistance and transport efficiency), mechanical strength, storage of water (capacitance) and starch, minimum hydrostatic pressures (Pmin), and proportions of fibers, vessels, and parenchyma, which were treated as a latent variable representing “cellular trade-offs.” We found that xylem functions (transport, mechanical support, water storage, and starch storage) were independent, a result driven by Pmin. Pmin was strongly and directly or indirectly associated with all xylem functions as a hub trait. More negative Pmin was associated with increased embolism resistance and tissue strength and reduced capacitance and starch storage. We found strong support for a trade-off between embolism resistance and transport efficiency. Tissue strength was not directly associated with embolism resistance or transport efficiency, and any associations were indirect involving Pmin. With Pmin removed from the model, cellular trade-offs were central and related to all other traits. We conclude that xylem traits are broadly governed by functional trade-offs and that the Pmin experienced by plants in the field exerts a strong influence over these relationships. Angiosperm xylem contains different cell types that contribute to different functions and that underpin trade-offs.

It is well established that transpiration and photosynthetic rates generally increase in resprouting shoots after fire in chaparral shrublands. By contrast, little is known about how plant hydraulic function varies during this same recovery period. We hypothesized that vascular traits, both functional and structural, would also shift in order to support this heightened level of gas exchange and growth. We examined stem xylem-specific hydraulic conductivity (Ks) and resistance to cavitation (P50) for eight chaparral shrub species as well as several potential xylem structural determinants of hydraulic function and compared established unburned plants and co-occurring post-fire resprouting plants. Unburned plants were generally more resistant to cavitation than resprouting plants, but the two groups did not differ in Ks. Resprouting plants had altered vessel structure compared to unburned plants, with resprouting plants having both wider diameter vessels and higher inter-vessel pit density. For biomechanics, unburned plants had both stronger and denser stem xylem tissue than resprouting plants. Shifts in hydraulic structure and function resulted in resprouting plants being more vulnerable to dehydration. The interaction between time since disturbance (i.e. resprouting versus established stands) and drought may complicate attempts to predict mortality risk of resprouting plants.

We study an optimization problem for a model of steady state water transport through plants that maximizes water flow subject to the constraints on hydraulic conductance due to vulnerability to embolism (air blockage of conduits). The model has an elementary geometric interpretation, and exhibits bottleneck behavior where one of the plant segments limits the overall optimal flow, sometimes in a counterintuitive way. The results show good agreement with experimental measurements and provide support for the hypothesis that leaves serve as a safety buffer protecting stems against excessive embolism., Comment: 13 pages, 2 tables, 3 figures

Resistance of stem xylem to water stress-induced cavitation and embolism among chaparral shrub species in California has been extensively studied, providing the opportunity to examine broad patterns in cavitation resistance. We used previously published as well as unpublished vulnerability to cavitation curve data from 16 chaparral shrub species of southern California to examine the variability of cavitation resistance across sites, regions, and seasons. Additionally, these data provided a unique opportunity to address a recent methodological debate within the field of plant hydraulics. We found that different methods, specifically a centrifuge method and a dehydration method, produced similar results (P = 0.184). Vulnerability to cavitation varied seasonally, with species exhibiting greater susceptibility to water-stress induced cavitation during the wet season (P = 0.003). Cavitation resistance did not differ among sites that were less than 10 km apart even though these sites differed in th...

It is well established that transpiration and photosynthetic rates generally increase in resprouting shoots after fire in chaparral shrublands. By contrast, little is known about how plant hydraulic function varies during this same recovery period. We hypothesized that vascular traits, both functional and structural, would also shift in order to support this heightened level of gas exchange and growth. We examined stem xylem-specific hydraulic conductivity (K

Vulnerability to cavitation curves describe the decrease in xylem hydraulic conductivity as xylem pressure declines. Several techniques for constructing vulnerability curves use centrifugal force to induce negative xylem pressure in stem or root segments. Centrifuge vulnerability curves constructed for long-vesselled species have been hypothesised to overestimate xylem vulnerability to cavitation due to increased vulnerability of vessels cut open at stem ends that extend to the middle or entirely through segments. We tested two key predictions of this hypothesis: (i) centrifugation induces greater embolism than dehydration in long-vesselled species, and (ii) the proportion of open vessels changes centrifuge vulnerability curves. Centrifuge and dehydration vulnerability curves were compared for a long- and short-vesselled species. The effect of open vessels was tested in four species by comparing centrifuge vulnerability curves for stems of two lengths. Centrifuge and dehydration vulnerability curves agreed well for the long- and short-vesselled species. Centrifuge vulnerability curves constructed using two stem lengths were similar. Also, the distribution of embolism along the length of centrifuged stems matched the theoretical pressure profile induced by centrifugation. We conclude that vulnerability to cavitation can be accurately characterised with vulnerability curves constructed using a centrifuge technique, even in long-vesselled species.

Lygodium microphyllum (Cav.) R. Br., a climbing fern native to the Pantropics of the Old World, is aggressively colonizing natural ecosystems in the Florida Peninsula. Here, we examined soil factors that might affect the fern’s invasiveness, specifically addressing the hypothesis that a release from natural belowground enemies contributes to its vigorous growth in Florida. We also investigated phenotypic differences of sporophytes raised from spores collected in Florida and the fern’s native range in Australia, hypothesizing that the Florida population would possess traits resulting in faster growth and superior competitive ability than the two Australian populations. We tested our hypotheses in parallel greenhouse experiments—one in Australia using soil from the fern’s native habitat, and another in Florida, USA, with soil from a recently colonized ecosystem. Fern growth rate and its principal determinants were expressed relative to the optimal growth with a common sand culture in each experiment and compared among treatments in which soil was altered through either sterilization or nutrient amendment, or both. Contrary to the expectation, the optimal growth rates in the sand culture were higher for Australian populations than the Florida population, while the comparatively poor growth of all populations in unaltered soil was stimulated by nutrient amendment and sterilization. The overall effect of sterilization, however, was muted under high-nutrient conditions, suggesting that the effect of soil sterilization may be due to greater nutrient availability in sterilized soils. The only exception was the local population from the site where the soil was collected for the experiment in Australia, which grew significantly faster in sterilized than in non-sterilized soil, and also more rapidly in response to soil insecticide application. Our results indicate that the invasiveness of L. microphyllum in Florida is not a simple phenotypic difference in inherent growth rate as predicted by the evolution of increased competitive ability hypothesis, but it may be mediated in part by release from soil-borne enemies that vary in their effectiveness even within the native geographical range of the fern.

We evaluated whether two indices of light availability resolved differences among microsites within deeply shaded understories (

Lianas are an important component of tropical forests, contributing up to 25 % of the woody stems and 35 % of woody species diversity. Lianas invest less in structural support but more in leaves compared to trees of similar biomass. These physiological and morphological differences suggest that lianas may interact with neighboring plants in ways that are different from similarly sized trees. However, the vast majority of past liana competition studies have failed to identify the unique competitive effects of lianas by controlling for the amount of biomass removed. We assessed liana competition in the forest understory over the course of 3 years by removing liana biomass and an equal amount of tree biomass in 40 plots at 10 sites in a secondary tropical moist forest in central Panama. We found that growth of understory trees and lianas, as well as planted seedlings, was limited due to competitive effects from both lianas and trees, though the competitive impacts varied by species, season, and size of neighbors. The removal of trees resulted in greater survival of planted seedlings compared to the removal of lianas, apparently related to a greater release from competition for light. In contrast, lianas had a species-specific negative effect on drought-tolerant Dipteryx oleifera seedlings during the dry season, potentially due to competition for water. We conclude that, at local scales, lianas and trees have unique and differential effects on understory dynamics, with lianas potentially competing more strongly during the dry season, and trees competing more strongly for light.

Shade-tolerant understory shrubs and subcanopy trees constitute most of the woody species in Neotropical moist forest, but studies demonstrating physiological differences among these species are few. Shade-tolerant species that coexist in the forest understory exhibit differences in leaf life span that have been associated with variation in physiological traits. We hypothesized that water relations of understory species with widely divergent leaf life spans differ in response to drought. Although severe drought is infrequent in Neotropical moist forest, we studied the water relations of shade-tolerant understory species with short or long leaf life spans during the severe 1991-1992 dry season on Barro Colorado Island, Panama. The predawn leaf water potential declined to -2.8 and -3.6 MPa during the dry season in Hybanthus prunifolius and Psychotria horizontalis, respectively, two species with short leaf life spans, but remained above -1.3 MPa in two species with long leaf life spans, Swartzia simplex and Ouratea lucens. The midday leaf water potential dropped as low as -3.4 and -4.5 MPa for H. prunifolius and 1? horizontalis, respectively. The osmotic potential of H. prunifolius and 1? horizontalis and another species with short leaf life span, Alseis blackiana, decreased early in the dry season, a period during which all three had substantially negative predawn water potential. In contrast, the osmotic potential of S. Simplex, 0. lucens, and Licania platypus, a third species with long leaf life span, declined late in the dry season, even though we observed little change in predawn water potential for S. simplex and 0. lucens. We conclude that the variable and potentially severe dry season in Neotropical moist forest can be sufficiently intense to severely limit soil moisture availability for understory plants. H. prunifolius and 1? horizontalis tolerated dehydration, whereas S. simplex and 0. lucens postponed dehydration.

We examined postfire regeneration of chaparral shrubs during an intense drought. This study focused on the demography and physiology of shrub species that resprout from a basal lignotuber following fire. We found significant levels of resprout mortality when intense drought occurred in the year following fire during the period of shrub recovery. Three of the seven sampled resprouting species had the greatest or near greatest levels of mortality ever recorded when compared to previous studies. Most shrub mortality occurred during the drought after individuals had resprouted (i.e. individuals survived fire, resprouted and then subsequently died). Physiological measurements of species with high mortality suggested that resprout stems were highly embolized and xylem hydraulic conductivities were close to zero during the peak of the drought. In addition, lignotubers of two of the three species experiencing high mortality were depleted of starch. Population densities of most shrub species declined after the drought compared with their prefire levels, with the exception of one drought tolerant obligate seeding species. Resprouting shrub species may deplete their carbohydrate reserves during the resprouting process, making them particularly vulnerable to drought because of the need to transpire water to acquire the CO2 that is used to supply energy to a large respiring root system. Drought appears to interact with fire by altering postfire shrub recovery and altering species abundances and composition of chaparral communities.

 Premise of the study: Vessels are the chief conduit for long-distance water transport in the majority of flplants. Vessel length is a key trait that determines plant hydraulic effi ciency and safety, yet relatively little is known about this xylem feature.  Methods: We used previously published studies to generate a new global data set of vessel length in woody plants. These data were used to examine how evolutionary history, plant habit, environment, and growth ring porosity infl uenced vessel length. We also examined the relationship between mean vessel length and mean vessel diameter and maximum vessel length.  Key results: Data on mean vessel length were available for stems of 130 species and on maximum vessel length for stems of 91 species. A phylogenetic analysis indicated that vessel length did not exhibit signifi cant phylogenetic signal. Liana species had longer vessel lengths than in tree or shrub species. Vessel diameter was not predictive of mean vessel length, but maximum vessel length strongly predicted mean vessel length. Vessel length did not vary between species that differed in growth ring porosity.  Conclusions: Many traits often assumed to be linked to vessel length, including growth ring porosity and vessel diameter, are not associated with vessel length when compared interspecifi cally. Sampling for vessel length has been nonrandom, e.g., there are virtually no data available for roots, and sampling for environment has been confounded with sampling for habit. Increased knowledge of vessel length is key to understanding the structure and function of the plant hydraulic pathway.

Lianas (woody vines) reduce growth and survival of host trees in both temperate and tropical forests; however, the relative strength of liana-tree competition in comparison to tree-tree competition remains unexplored. When controlling for biomass, lianas may have greater competitive effects than trees because the unique morphology of lianas allows them to reach the forest canopy at relatively small stem diameters and deploy a substantial crown above their host. We tested the hypothesis that lianas have a greater negative effect on canopy trees than do trees of similar biomass with a liana- and tree sapling-cutting experiment in a seasonal tropical moist forest in Panama. The response of canopy trees to the cutting treatments was characterized as the change in their daily water use by measuring their sap velocity before and after cutting. We compared the responses of canopy trees around which a similar biomass of either lianas or tree saplings had been cut to control trees with no cutting. Liana cutting increased canopy-tree sap velocity by ∼8% from before to after cutting relative to control trees during the dry season. In contrast, canopy-tree sap velocity did not respond to tree cutting, probably because trees with biomass similar to lianas were confined to the forest understory. We observed a similar pattern of sap velocity changes during the wet season, but treatment differences were not significant. Our results demonstrate that release from liana competition, but not tree competition, resulted in increased water transport in canopy trees, and suggests that relative to their biomass, lianas have greater competitive effects on canopy tree performance than do competing trees.